Handbook of Waterfowl Behavior, by Paul JohnsgardCopyright (c) 2014 University of Nebraska - Lincoln All rights reserved.http://digitalcommons.unl.edu/bioscihandwaterfowl
Recent documents in Handbook of Waterfowl Behavior, by Paul Johnsgarden-usThu, 03 Jul 2014 17:45:46 PDT3600Handbook of Waterfowl Behavior: Indexeshttp://digitalcommons.unl.edu/bioscihandwaterfowl/22
http://digitalcommons.unl.edu/bioscihandwaterfowl/22Thu, 05 Jun 2008 07:52:57 PDT
General index: The principal page reference for each species or group is indicated by boldface type. Subordinate page references for each species are listed only under the standardized English names used in this book, as shown in the Appendix. Commonly used alternate names, or names referring to certain races or species groups, are included in the index but only principal page references are listed under these names. Page references to illustrations of species or of specific displays are indicated by italics. The Appendix and Summary have not been indexed.

Scientific Names: This index provides a reference only to the page where the principal discussion of each genus or species begins. The general index should be consulted for locations of illustrations and for subordinate references. Several commonly used generic names, here considered synonyms or subgenera, are included for cross reference and are set off by quotation marks.
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Paul A. JohnsgardHandbook of Waterfowl Behavior: Works Citedhttp://digitalcommons.unl.edu/bioscihandwaterfowl/21
http://digitalcommons.unl.edu/bioscihandwaterfowl/21Thu, 05 Jun 2008 07:50:38 PDT
Approximately 150 works cited. (7 pages)
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Paul A. JohnsgardHandbook of Waterfowl Behavior: Summary and Synopsis of the Family Anatidaehttp://digitalcommons.unl.edu/bioscihandwaterfowl/20
http://digitalcommons.unl.edu/bioscihandwaterfowl/20Thu, 05 Jun 2008 07:49:22 PDT
The major sexual behavior patterns of the species of Anatidae may now be reviewed and summarized, for the purpose of clarifying evolutionary relationships in the family, tracing the evolution of behavior patterns through various taxa, and evaluating the importance of certain behavior patterns as potential isolating mechanisms.
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Paul A. JohnsgardHandbook of Waterfowl Behavior: Tribe Oxyurini (Stiff-tailed Ducks)http://digitalcommons.unl.edu/bioscihandwaterfowl/19
http://digitalcommons.unl.edu/bioscihandwaterfowl/19Thu, 05 Jun 2008 07:46:00 PDT
The stiff-tailed ducks constitute a unique section of the Anatidae that is possibly the most isolated of all the tribes with the exception of the Anseranatini. There are eight species which almost certainly belong in the group, plus one more that is only very tentatively included. The tribe is of worldwide occurrence. Seven of the species have long, narrow, and stiffened tail feathers that function as rudders in underwater swimming, at which all species are very adept. These species also have a dense and shiny body plumage much like that of grebes, but lack metallic coloration altogether. The typical species have short, thick necks with loose-fitting skin that can be expanded through the inflation of the esophagus or special air sacs. All species have large feet and their legs are placed well toward the rear, which results in a poor walking ability. All species but the most aberrant one (the white-backed duck) exhibit sexual dimorphism. Vocalizations are extremely variable, and male calls are often produced by extra-tracheal structures. Sexual maturity is probably reached in the first year in all but one of the species (the musk duck), and pairs are probably renewed yearly in most and quite possibly all species. Male displays are generally elaborate and tend to produce sound. Nests are generally built over the water surface and the eggs are generally chalky white and relatively large, the young being very precocial at hatching.
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Paul A. JohnsgardHandbook of Waterfowl Behavior: Tribe Mergini (Sea Ducks)http://digitalcommons.unl.edu/bioscihandwaterfowl/18
http://digitalcommons.unl.edu/bioscihandwaterfowl/18Thu, 05 Jun 2008 07:45:58 PDT
The tribe includes 20 species of which two are extinct. Except for two Southern Hemisphere species, the group is of northern and arctic distribution. All species consume a high proportion of animal material, although they are by no means all "sea" ducks. All dive extremely well, although the manner of diving varies in that some species usually open their wings when diving and others never do. All the species except the isolated Southern Hemisphere forms exhibit considerable sexual dimorphism in plumage, voice, and behavior, and there is much sympatry of ranges. Some genera (Somateria, Polysticta, Clangula, Melanitta) normally nest on the ground, and those species which usually nest in the open (the eiders) tend to have cryptic and disruptively patterned female plumages. The other species tend to nest in crevices or holes in the ground (the harlequin), or in trees, and females of these species lack such cryptic coloration. Metallic coloration is restricted to male head plumage in most species, but there is a metallic-colored speculum in two genera (Polysticta and Histrionicus). The other species either have white wing-specula or lack any wing patterning.
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Paul A. JohnsgardHandbook of Waterfowl Behavior: Tribe Aythyini (Pochards)http://digitalcommons.unl.edu/bioscihandwaterfowl/17
http://digitalcommons.unl.edu/bioscihandwaterfowl/17Thu, 05 Jun 2008 07:43:53 PDT
The dabbling ducks grade almost imperceptibly into the pochard group considered here, and whether Marmaronetta and Rhodonessa should be included in one tribe or the other may be open to some question. As here constituted, the tribe includes 16 species of almost world-wide distribution which differ from the preceding group in several minor details. The hind toe has a larger lobe than occurs in the dabbling ducks, and it presumably aids in diving. The feet are also correspondingly larger and are set farther apart and more to the rear than in Anas. One of the most clear-cut differences is in the tracheal structure of the male. Instead of a rounded and entirely osseous tracheal bulla such as occurs in Anas, the bulla is larger, rather angular, and contains several membranceous fenestrae of varying sizes. Although the bulla structure of Netta and Aythya is strikingly different from that of the Anas, the genera Marmaronetta and Rhodonessa provide such perfect intermediate stages that the evolutionary development of this structure is very clear. In addition, the tracheal tube of males of nearly all species (Aythya novae-seelandiae and A. collaris are apparently exceptions) varies in diameter and is enlarged toward the middle. Unlike the typical dabbling ducks, none of these species have metallic-colored specula, and metallic coloration is restricted to the head plumage of the males of some species. Although some species (especially Rhodonessa and Netta) frequently feed on the surface, all species dive well and typically do not open their wings when submerging. Most species are predominantly vegetarian, but the scauplike ducks tend to consume a high proportion of animal material. Nests may be built on land, usually near water, or on reed beds over the water surface, and females lack the disruptively marked plumage pattern typical of female dabbling ducks. Downy young tend to lack eye stripes and cheek marks, and also generally have reduced back spotting. Most if not all species become sexually mature during the first year; the scauplike ducks may take slightly longer. All species exhibit sexual dimorphism in plumage and/or soft- art coloration, and nearly all species have distinct eclipse plumages. Although metallic-colored specula are absent, the secondaries have a contrasting white pattern in many species.
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Paul A. JohnsgardHandbook of Waterfowl Behavior: Tribe Anatini (Surface-feeding Ducks)http://digitalcommons.unl.edu/bioscihandwaterfowl/16
http://digitalcommons.unl.edu/bioscihandwaterfowl/16Thu, 05 Jun 2008 07:42:29 PDT
The tribe of surface-feeding, or dabbling, ducks is the largest single tribe in the family. There are 40 species in the tribe as it is constituted here. Contrary to the arrangement of Delacour (1956), the ringed teal is included in the Cairinini; and the crested duck, included in the Tadornini by Delacour, is here considered a typical dabbling duck. In addition, the pink-headed duck has been removed from the Anatini and placed in the Aythyini with the pochards, and the freckled duck has been removed from the tribe and is considered a primitive species having anserine relationships. The marbled teal has been removed from the genus Anas and placed in a monotypic. genus which, I believe, provides an evolutionary link between the dabbling ducks and the pochards.

The Anatini are world-wide in distribution and include the most numerous and widespread species of the family. Most species (and especially those of the shoveler group) forage on the water surface, gathering food from the surface by up-ending or "tipping-up," or, more rarely, by diving. Most species open their wings when diving. A few species, such as the wigeon, graze, and nearly all species are predominantly vegetarians. Nearly all nest on the ground, but a few species nest in holes or crevices. The downy young are much like those of the perching ducks and are usually strongly marked with brown and white or with yellow. All species mature their first year, and pair bonds are generally renewed yearly. Most species exhibit sexual dimorphism in plumage, and metallic coloration is usually restricted to the head and speculum. The males of most sexually dimorphic species have a distinct eclipse plumage. Nearly all species have metallically colored wing specula. Males of all species studied to date except one have tracheal bullae which are ossified throughout; the marbled teal is the exception, and its bulla has numerous membranaceous fenestrae similar to those found in the pochard group.
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Paul A. JohnsgardHandbook of Waterfowl Behavior: Tribe Cairinini (Perching Ducks)http://digitalcommons.unl.edu/bioscihandwaterfowl/15
http://digitalcommons.unl.edu/bioscihandwaterfowl/15Thu, 05 Jun 2008 07:41:05 PDT
Delacour and Mayr (1945) placed the perching duck tribe after the pochards (Aythyini) and adjacent to the sea ducks (Mergini), but hybridization evidence (Johnsgard, 1960a) clearly indicates that the group belongs between the shelducks and the dabbling ducks and closely adjacent to the latter. There is other evidence to support this arrangement. The tracheae of shelducks, perching ducks, and dabbling ducks are all very similar, possessing bullae which in most species are osseous and rounded, and lacking enlargements of the tracheal tube. The downy young of perching ducks and dabbling ducks are very similar, and are usually patterned with dark brown and white or yellow. Woolfenden (1 96 1) has advocated merging the perching ducks and dabbling ducks into a single tribe (Anatini), but although the two groups do tend to overlap in some of their characteristics, I believe that merging them would result in an unduly large and heterogeneous tribe. Such action would, however, resolve the problem of the tribal allocations of such species as the Brazilian teal and ringed teal. Some species of perching ducks mature in two years, although the smaller species mature their first year. Nearly all perching ducks have metallic coloration on the upper-wing surface. With few exceptions they do not dive well or frequently, and apparently all use their wings when submerging. Most of them dabble for their food, and some also graze. Nearly all species are hole-nesters, and have relatively long incubation periods. As here constituted the tribe contains thirteen species; those included in it by Delacour (1959), plus an additional species, the ringed teal (Anas leucophrys of Delacour), which is here considered to comprise a separate genus (Callonetta).
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Paul A. JohnsgardHandbook of Waterfowl Behavior: Tribe Tachyerini (Steamer Ducks)http://digitalcommons.unl.edu/bioscihandwaterfowl/14
http://digitalcommons.unl.edu/bioscihandwaterfowl/14Thu, 05 Jun 2008 07:39:41 PDT
I consider the three species of steamer ducks of the genus Tachyeres to constitute a separate tribe. These authorities agree that although steamer ducks are perhaps most closely related to the true shelducks, the steamers are sufficiently different to warrant removing them from the shelduck tribe. Their downy young lack the strongly contrasting coloration typical of shelducks, and the adult plumage pattern is distinct both from that of the shelducks and from that of the other anatine groups. There is a slight sexual dimorphism in bill color, head color, and possibly in the degree of tail-feather curling. Pair bonds appear to be strong, and Murphy (1936) was of the opinion that steamer ducks pair for life. At least two years are required for the birds to reach maturity. There are apparently three similar-appearing species, of which two are virtually flightless. The voices of the sexes are very different, and the males have tracheae with rounded, osseous bullae which are similar to those of Anas. There is no metallic coloration in either sex; the wings have white secondaries and secondary coverts. All species dive extremely well, and the birds feed to a great extent on marine animal life off coastal South America and the Falkland Islands.
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Paul A. JohnsgardHandbook of Waterfowl Behavior: Tribe Tadornini (Sheldgeese and Shelducks)http://digitalcommons.unl.edu/bioscihandwaterfowl/13
http://digitalcommons.unl.edu/bioscihandwaterfowl/13Thu, 05 Jun 2008 07:39:31 PDT
The tribe includes the 16 species of sheldgeese and shelducks ("Lophonetta" and Tachyeres are here excluded from the tribe) which provide such a smooth transition from the true geese to the true ducks that it is difficult to establish a dividing line between the two groups. I believe that the line is best placed between Cereopsis on the anserine side and Cyanochen on the anatine side. Cyanochen is clearly a sheldgoose, but the plumages and voices of the sexes are practically alike, as are also their displays. Cyanochen and the other Tadornini are characterized by a downy plumage that is strongly marked with dark and white, by a tendency to nest in cavities, and by the fact that they forage either by grazing (sheldgeese) or by grazing and dabbling (shelducks). Metallic plumage coloration is encountered in this group, as are vermiculated feather patterns; both are typical of most of the rest of the Anatinae. Pair bonds appear to be fairly strong in sheldgeese, but less strong in shelducks. Unlike that of the true geese, pair-forming, or courtship, behavior is conspicuous, and in this group may be observed the basic pair-forming mechanism which is to be found, in varying degrees of refinement, in all other tribes of Anatinae. Simply stated, this is the tendency of the female to Incite (Lorenz, 1951-1953) males to attack other males or females, and to "select" her mate on the basis of the male's reaction to this Inciting. Here sexual selection enters the picture, and the complex male plumage patterns and male courtship- behavior patterns can be understood and interpreted only by reference to the principle of sexual selection.
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Paul A. JohnsgardHandbook of Waterfowl Behavior: Tribe Stictonettini (Freckled Duck)http://digitalcommons.unl.edu/bioscihandwaterfowl/12
http://digitalcommons.unl.edu/bioscihandwaterfowl/12Thu, 05 Jun 2008 07:37:45 PDT
I include the Australian freckled duck, as the only member of a proposed tribe Stictonettini, as I originally suggested in 1960 on a tentative basis. Previously the freckled duck had been generally considered an aberrant dabbling duck with several primitive features, such as a reticulated tarsus, simple tracheal structure (illustrated in Johnsgard, 1961c), and absence of metallic plumage coloration or distinct plumage dimorphism. No hybrids have been recorded involving this species, which occurs widely but infrequently over the southern half of Australia.
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Paul A. JohnsgardHandbook of Waterfowl Behavior: Tribe Anserini (Swans and True Geese)http://digitalcommons.unl.edu/bioscihandwaterfowl/11
http://digitalcommons.unl.edu/bioscihandwaterfowl/11Thu, 05 Jun 2008 07:36:30 PDT
The swans and true geese of the present tribe include about 21 species which are primarily temperate and arctic in distribution, with most forms occurring in the Northern Hemisphere. They are generally large grazing or dabbling birds, and all are highly social and have strong family bonds. Plumage patterns are the same in both sexes and tend to be relatively simple, with visual signal characteristics generally restricted to the bill, head, and rump. For the most part the species are very vocal, and in the true geese vocalizations probably reach the highest degree of development in the entire family.
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Paul A. JohnsgardHandbook of Waterfowl Behavior: Tribe Dendrocygnini (Whistling Ducks)http://digitalcommons.unl.edu/bioscihandwaterfowl/10
http://digitalcommons.unl.edu/bioscihandwaterfowl/10Thu, 05 Jun 2008 07:35:12 PDT
The whistling, or "tree," ducks comprise a group of eight species of world-wide, though primarily tropical, distribution. A single genus, Dendrocygna, is generally accepted, and indeed the species form such a homogeneous group that it is extremely difficult to establish intrageneric differences upon which to judge probable species relationships. In most respects, whistling ducks are very gooselike, differing from geese mainly in their more specialized tracheal structure, their whistling voices, and their distinctive downy plumage patterns. They possess relatively long legs and large feet, and they can dive remarkably well.
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Paul A. JohnsgardHandbook of Waterfowl Behavior: Tribe Anseranatini (Magpie Goose)http://digitalcommons.unl.edu/bioscihandwaterfowl/9
http://digitalcommons.unl.edu/bioscihandwaterfowl/9Thu, 05 Jun 2008 07:33:59 PDT
The first subfamily and tribe of the Anatidae consists of the single genus and species Anseranas semipalmata, or magpie goose. This species deviates in almost every respect from the rest of the Anatidae, and anatomical evidence indicates that it has some affinities with the South American family of screamers, the Anhimidae (Delacour, 1954; Woolfenden, 1961). The screamers are a group of semiterrestrial birds which in turn have some of the characteristics of gallinaceous birds. It appears that the screamers and the magpie goose represent major stages in the evolution of the typical waterfowl from a gallinaceous ancestor. Besides the numerous anatomical similarities between the magpie goose and the screamers, a few other, obvious similarities are of interest.
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Paul A. JohnsgardHandbook of Waterfowl Behavior: Frontmatter and Introductionhttp://digitalcommons.unl.edu/bioscihandwaterfowl/8
http://digitalcommons.unl.edu/bioscihandwaterfowl/8Thu, 05 Jun 2008 07:33:57 PDT
Acknowledgments Contents List of Illustrations Background and Objectives of This Study Value and Limitations of Behavior as a Taxonomic Tool Biological Characteristics of the Family Anatidae
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Paul A. JohnsgardHandbook of Waterfowl Behaviorhttp://digitalcommons.unl.edu/bioscihandwaterfowl/7
http://digitalcommons.unl.edu/bioscihandwaterfowl/7Thu, 05 Jun 2008 07:31:47 PDT
The present study was undertaken to test and evaluate the various taxonomic arrangements of the family, to discover some of the trends of behavioral evolution, and to provide a basis for future workers to use in naming, describing, and evaluating the behavioral patterns observed in waterfowl.

The major sexual behavior patterns of the species of Anatidae are reviewed and summarized, for the purpose of clarifying evolutionary relationships in the family, tracing the evolution of behavior patterns through various taxa, and evaluating the importance of certain behavior patterns as potential isolating mechanisms.